A brief history of the scientific study of tropical African inland waters

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Authors Talling, Jack

Download date 06/10/2021 16:06:49

Link to Item http://hdl.handle.net/1834/22213 TROPICAL AFRICAN INLAND WATERS 3

A BRIEF HISTORY OF THE SCIENTIFIC STUDY OF TROPICAL AFRICAN INLAND WATERS

JACK TALLING

Dr J.F. Talling, Freshwater Biological Association, Far Sawrey, Ambleside, Cumbria LA22 0LP, UK

Introduction The inland waters of Africa, and especially its large rivers, determined much of the original geographical exploration of the continent. Subsequently, their scientific study has contributed to the science of inland waters in general and of tropical inland waters in particular. The history of this tropical study is outlined below. It draws upon personal experience in several African universities and a research institute, vivid impressions of aquatic habitats and communities within African landscapes, and a desire to move from individual descriptions to wider comparisons and the underlying fundamentals. In any history of scientific endeavour in Africa, the human factor deserves special attention. In colonial times much science, like trade, followed the flag. In Africa, unlike South America, this phase extended over – and benefited from – a period of rapid growth in freshwater science. Later, after the 1950s and widespread political independence, the science continued from both pre-established and new institutions – mainly universities and fishery institutes. These gave new opportunities to indigenous staff, and often hosted relatively brief expeditions from abroad. International and inter-governmental projects grew in importance. They were planned with regard to perceived needs, technical and socio- economic, of a region; but they could also enrich fundamental insights within the subject. The wide coverage has necessarily involved the exclusion of detail on the contributions of individuals, although examples appear in the numerous references. In this I follow the pattern adopted in a history of British contributions to freshwater science244.

The situation in 1900 By this date the disposition of the main rivers and lakes (Fig. 1) was reasonably well known. It was evident that most of the rivers had a strongly variable – indeed rhythmic – discharge, linked to the alternation of rainy and dry seasons. That of the Nile had been followed, by water level in Egypt, from ancient times; the crucial connection of its floodwater to

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right)

) crustal warping FIG. 1. Map of Africa with named waterbodies (shallow lakes underlined)243.

top left rains in Ethiopia was proposed in 1668 by the Jesuit Father Lobo236. Lakes (Fig. 2) were numerous, some very large and influenced by large-scale

tectonic events like crustal warping (e.g. Victoria, Chad) and rifting (e.g. ( ) rifting – Lake Albert;

Tanganyika). Systems of both open and closed drainage were involved, Ethiopia. Lake Aranguadi, with a corresponding wide range of water salinity. Only in a few cases were there analyses of the chemical composition, such as that obtained by

Henry Wellcome – subsequently an influential figure in tropical medicine bottom left – for a salt lake in western Uganda207.

A beginning had been made towards assessment of aquatic faunas and floras. Most collections were sporadic, as of an early sample of algae from 57 Lake Nyasa . Near the end of the century there were two examples of more sustained enquiry. The aquatic fauna of Lake Tanganyika had – Lake Victoria; ( Victoria; – Lake volcanic explosion crater – attracted attention by the presence of endemics and the resemblance of lakes of varied origin: ( FIG. 2. African some species – especially thick-shelled molluscs – to marine forms. This

Freshwater Forum 26 (2006) Freshwater Forum 26 (2006) 6 JACK TALLING TROPICAL AFRICAN INLAND WATERS 7 was taken by Moore as evidence of a direct marine origin for the lake, 192 leading to his expeditions of 1895 and 1898 and subsequent book that suggested derivation from a Jurassic sea. The second example concerned the water-mass and plankton of Lake Nyasa (later Malawi), investigated during a German expedition of 1899–1900. Using a simple sampler, the expedition laboriously discovered the first thermocline to be described from a tropical lake82. Net-samples of plankton, gathered throughout 1899 and later examined by Schmidle, were used to construct a first rudimentary 234 account of the seasonal changes of phytoplankton in a tropical lake .

The period 1900–1925 This period saw some notable acquisitions of information regarding aquatic environments and their biota, although integrated ecological science had not yet arrived. The vertical configurations (bathymetry) of various large lakes, including Victoria and Tanganyika (Fig. 1), were mapped and found to range widely. The work of Stappers in 1911–13 on 237 L. Tanganyika disclosed a considerable volume of water at depths exceeding 1000 m. Vertical relationships were also important in the regulation of flow in rivers. Thus a ‘stepped’ longitudinal section of the 167 Nile included Lake Plateau, Rift Valley, savanna, Sudd swamp and desert cataracts (examples in Fig. 3).

Various expeditions and their collections made somewhat disconnected contributions to aquatic science. Examples relating to water chemistry are cited by Talling & Talling246 and others to freshwater algae by Brook et al.30. Zoological work is exemplified by the plankton collections examined by Daday48 from Lake Victoria. There was the remarkable discovery of aquatic animals from within the Sahara212. Three examples of more sustained and systematic study are of note. Shortly after 1900 the Wellcome Laboratories were set up at Khartoum in 10 FIG. 3. Dissimilar river sectors down the Gordon Memorial College; from them Beam made chemical analyses the Nile: (top left) savanna, southern over several years of water from the Blue and White Niles that Sudan; (bottom left) desert cataract, demonstrated considerable seasonal and longitudinal changes. Collections Nubia; (above right) floodplain swamps of fishes from the Congo basin and the Nile were available in Europe and (‘Sudd’). 26, used by Boulenger to produce major monographs on the two fish faunas 27. Lastly, the controversial ‘Tanganyika Problem’ of Moore was tested further by the expedition of Cunnington (1904–05). From its collections and with the cooperation of many group-specialists, Cunnington assembled a ‘comparative limnology’47 and rejected the original hypothesis of a direct marine origin – already doubted from other work on the gastropod molluscs213.

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The beginnings of integrated ecology: the years 1925–1945 differences that were dominated by a wide range of titration alkalinity due to bicarbonate and carbonate130. The biota were often very dense, and The following 20 years were a period of advancing colonial administration involved a carbon flow that Jenkin investigated by experiments on in many African territories, in which research was stimulated by economic photosynthesis in situ (adapted from the then recent work of Marshall and and social issues. Two major areas were hydrology and fisheries. The Orr in Scotland) and observations on the filter-feeding mechanism of former involved estimation of water budgets and the flow gauging of many flamingos131. A second and larger Cambridge expedition, in 1930–31, was rivers. One was the Nile, on which the original Aswan Dam (Fig. 4) was led by Worthington – clearly enthused by his earlier African experience on created, and for which Hurst and his associates in the Physical Department lakes Victoria and Albert. It too dealt comparatively with a range of lakes, of the Ministry of Public Works at Cairo gathered information from and with communities from phytoplankton to fishes. One participant, sources to delta in a series of hydrological monographs The Nile Basin 124 Beadle, provided a wide range of information on water chemistry and (e.g. ). The broad picture for the Nile was later summarised in a more 6 123 functional relationships which included photosynthesis-depth distribution popular book by Hurst , and for the hydrology of tropical Africa in 5 271 2 and ecology of swamp invertebrates . Besides zooplankton studies , general by Balek . Worthington took up the comparative faunistics and biology of the fish Initiatives that concerned inland fisheries had a greater influence on the communities, and subsequently suggested interpretations of their development of aquatic ecology. The most significant began with the 269 92 evolutionary differentiation in Africa – a topic that later attracted many fisheries survey of Lake Victoria by Graham and Worthington in 1927– biologists76, 79, 95, 218. Post-expedition he was helped in the assessment of 28. This extended beyond fish faunistics, distribution and abundance to zooplankton abundance by a student, Kate Ricardo271, who later was to problems of changing thermal stratification267 and day-night (diel) vertical 268 contribute in the field with investigation of fish communities in several migration of zooplankton . All these subjects were taken up further in large rift-lakes of Central Africa18, 224. work on the nearby Lake Albert. The investigation of diel change in Until the 1930s there was but little knowledge of physical and chemical Victoria was the first such work for tropical fresh waters. aspects of vertical stratification in the deeper rift lakes. By 1913, Stappers In the years immediately following, there were two further British and others had shown that the deepest lake of all – Tanganyika – was expeditions that closely interrelated biological and environmental features vertically stratified. In 1935–36 another Belgian scientist, Damas, of East African lakes. That of Penelope Jenkin, in 1929, was a one-woman contributed a wider range of findings – physical, chemical and biological – expedition that developed from friendship with a member of the Leakey 51 166 on the stratification of lakes Kivu and Edward . He showed that family . It took in Kenyan lake waters lacking surface outflows, with deoxygenation of their deeper water was accompanied by large increments of ionic concentration, suggestive of long continuation rather than annual interruption of density stratification. Shortly afterwards, Beauchamp11 made comparable physical and chemical measurements on L. Tanganyika with a longitudinal sequence of stations – and a hand-operated winch! He demonstrated the existence of a large anoxic water-mass below 100–200 m depth. The work was later, in 1939–40, followed up by measurements on changes of stratification in Lake Nyasa (later Malawi)13. Here Beauchamp found short-period oscillations in the depth of individual isotherms that he interpreted as evidence for longitudinal internal waves or seiches, as well as some seasonality in vertical entrainment. Confirmatory evidence of these features was obtained by others in the 1950s and later decades24, 64 when longer time-series of observations were made, some linked with work on fisheries65. There were pioneer studies of the fish stocks by FIG. 4. Impoundments of man-made lakes: (left) Bertram (Ricardo) et al.18 and later by Lowe161. The last, and the work of original Aswan Dam, photo. 1955; (above right) Beauchamp and Worthington, were forerunners of much later involvement Kariba Dam, photo. 1983. in African freshwater research from the Freshwater Biological Association80 in Britain.

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In Ethiopia the beginnings of freshwater research were unusual. Before Corbet43, Fryer77, Garrod83, Hamblyn101, Elder and van Someren253, and 1935 only a few expeditions with collections of samples205 were recorded. subsequently Jackson127, Mann174 and Welcomme261. Later the brief period of Italian rule contained several surveys of major rift Another initiative within fisheries research for territories of Central lakes256 and the elevated Lake Tana193, as well as various small crater Africa was founded in 1951 from the Colonial Office in London. This, the lakes. Joint Fisheries Research Organization (JFRO), operated during 1951–63 To 1945 therefore, in tropical Africa overall, we can record important over a series of lakes in or bordering the then Northern Rhodesia and elements of progress with the compositional-structure and (in much lesser Nyasaland – Bangweulu, Malawi128, Kariba104 and Tanganyika44. Its degree) dynamics of freshwater environments and their biological scientific staff included Fryer, Harding, Iles, Jackson and later Bowmaker communities. In the biological sphere there were – too many to summarize and Coulter; Jackson has recently given a lively account of its history127. here – further acquisitions of taxonomic, faunistic and floristic information Although fish biology, taxonomy and fisheries received most attention, for aquatic organisms from plankton to fishes. River hydrology excepted, there were also important contributions in associated chemical surveys104, there was a marked deficiency of systematic observations maintained over lake hydrography44, plankton ecology44 and the general biology of aquatic long periods. invertebrates78. Some more local surveys of fisheries, also supported from the British Colonial Office, had productive collaborations with other 1945–1965: era of later colonial administration and its disappearance hydrobiologists. Examples include the work of Holden with Green on the River Sokoto117, 118 and with Green and Evans on Lake Albert70, 93, and that With the cessation of hostilities in 1945, initiatives for freshwater research on Lake Chad from fisheries-based projects in northern Nigeria119, 227. came from several European countries with colonial or dependent The post-1945 years also saw a major development of universities or territories in Africa. In the main they linked with the development of university colleges, at which the research attractions of African inland fisheries institutes and universities, and so with long-maintained waters were recognised and new members of staff with freshwater operations. expertise appeared. In Uganda, at Makerere College (later University), Nevertheless, expeditions continued to be made. Two from Belgium there were Beadle and Lind who had special interests in lakes8 and papyrus were important for exploration of the large rift lakes of Tanganyika (1946– swamps that included ecophysiology7 and community floristics157. 47) and Kivu, Edward and Albert (1952–54). Inter alia, they provided 110 36 Hartland-Rowe drew attention to a lunar rhythm in insect emergence . In bathymetric maps of detail and precision from the use of echo-sounding , the Sudan, at the University College (later University) of Khartoum, was structures of physical and chemical stratification37, 147, and comparative 252 151, 257 Rzóska (an émigré Polish hydrobiologist) and later Brook, Prowse, studies of chemical composition, plankton , zoobenthos and Thornton and myself. The College set up a Hydrobiological Research Unit macrophytes251. Neither was primarily devoted to problems of fisheries, 219, 220 258 in 1953, equipped with research launch (Fig. 5) and truck that enabled although information on fish faunas and their food accrued. some immense longitudinal surveys of the upper Nile system to be made20, The prospect of more productive fisheries in major African lakes led to 230, 241. The work included some pioneering estimations of photosynthetic the founding of several institutes and mobile teams. Of high scientific productivity223, but much concentrated on the distribution and seasonality distinction, and long persistence, was the East African Fisheries Research of plankton and associated water characteristics of the White and Blue Organisation (EAFRO) set up in 1947 beside Lake Victoria at Jinja, Niles near Khartoum. There, one predominant influence was the seasonal Uganda. It was then unusual (and sometimes criticised!) for the wide range retention of water in reservoirs upstream31, 223, 232. Another was the reverse of freshwater science that was pursued, partly reflecting the broad interests 12 wash-out in annual floodwater and reservoir discharge, as had been of its first Director, Beauchamp. His early Annual Reports included some followed by Abdin1 for phytoplankton of the Aswan reservoir in upper speculative portrayals of freshwater ecosystem science. In these early years Egypt. During the late 1950s there was a massive and troublesome the staff represented interests in tropical fish biology95, 163, water chemistry, 73 168 invasion of the White Nile by the floating water hyacinth Eichhornia hydrography and phytoplankton , and aquatic insects . The range of crassipes88 that was already established, and studied, in the Congo (Zaïre) research was enhanced then and later by encouragement of visiting 17, 228 39 system . The latter and the upper-middle Niger, waters with rich fish scientists, including Carter with work on swamp chemistry , myself in 221 22a, 49, 53, 175 246 faunas , were investigated by Belgian and French scientists . 1956 and with my wife in 1960–61 . The later staff members included In the 1950s there was a Belgian initiative, not directly connected with fisheries or universities of Africa, that had no equivalent elsewhere. A

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Of these, the events ensuing in Lake Kariba were among the most dramatic and best known. They were traced by a series of research groups that included universities at Salisbury (later Harare)171, 189 and Johannesburg (‘Wits’), organisations for fishery research (JFRO104, 127, Kariba Fisheries Institute178, F.A.O. Fisheries Institute4) and the Nuffield Lake Kariba Research Station at Sinamwenda29, 182. Early features included a transitory increase in concentration of plant nutrients (e.g. nitrate); deoxygenation with sulphide production in hypolimnetic water; the displacement of a rich bush fauna; the spread of a water-fern, Salvinia molesta, over a considerable part of the lake surface189; and the successful introduction from Lake Tanganyika of a small pelagic ‘sardine’ Limnothrissa miodon178 that later supported a vigorous fishery. Notable scientific contributions included the demonstration of successive mixing/stratification events in a chain of sub-basins4, the co-existence of small tidal and surface seiche oscillations in water level260, and the influence of diel vertical migrations for the pelagic food chain15. Long- term changes were documented and assembled by Coche, McLachlan and 183 FIG. 5. Research on the White Nile, Sudan, 1954: cabin of the launch Malakal, with Marshall, and placed in a generalised African setting . J. Rzóska (standing) and the author. Much less research effort was given to another man-made lake on the Zambezi, Cahora Bassa23. For this political and military hazards52 were hydrobiological station was established at Uvira near the northern end of largely responsible. Nevertheless there was a notable investigation of Lake Tanganyika. Its scientific workers included Dubois, Marlier and plankton dynamics by Gliwicz90. Favourable conditions with available Symoens. From here time-sequences of observations were possible, which expertise were available for work on Lake Nasser (Egypt) and lakes Volta contributed to knowledge of interlinked hydrography, water chemistry58, 239 176 and Kainji in West Africa. Both local and international resources were and planktonic production ; also of the benthic invertebrates of this involved. Thus, on the Nile from Aswan and Cairo, studies were made of most ancient and unique lake. the radically altered hydrological regime and its consequences for water Much further south, in the 1960s, another Hydrobiology Research Unit characteristics68, hydrobiology100 and fisheries148. Much of the research on was active in the University College of Rhodesia and Nyasaland (later Lake Volta was carried out by university-based workers from Ghana149 and University of Zimbabwe) at Salisbury (later Harare). Subsequent 177, 190 190 226 Britain. The Food and Agricultural Organisation (FAO) made a major participants, including Marshall , Mitchell and Robarts , contribution with the work of Petr and others on zoobenthos216 and fish explored conditions in a variety of reservoirs or ‘dams’ that included Lake biology215. This organisation also supported research on the smaller Lake McIlwaine (= Chivero). Later much attention shifted to the new man-made Kainji of the Niger19, 115, 136, with further continuation from Imevbore and Lake Kariba. others126 of the University of Ife and work on the main river up- and down- stream97. Also studied in Nigeria were the effects on phytoplankton of 1965–1985: development of large man-made lakes impoundment on a smaller river67. A reservoir in Tanzania was the subject and international projects of a many-sided investigation from Britain55. Within the new lakes time- In the years after 1960, large ‘man-made lakes’ 163, 202 were created in successions were evident in communities of plankton, benthos and fish. tropical Africa for water distribution and power generation. These altered Interesting interrelations emerged, some involving zoobenthos, river regimes, with over-year storage, and provided unique examples of periphyton/aufwuchs and plankton as alternative and changing sources of succession in aquatic environments and biota. Thus Lake Kariba on the food for fish217 – a group in which seasonal biological rhythms, developed Zambezi (Fig. 4) began to fill in 1958; there followed Lake Nubia-Nasser21, under riverine conditions, were liable to change adaptively with flow 148 on the Nile, Lake Volta in Ghana149, Lake Kainji in Nigeria126, and Lake variation126a. Cahora Bassa23 in Moçambique.

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Some hydrological projects involved canals rather than dams. The Coe42, Gaudet86, 87, Kalff134, Litterick181, Lind156, Mavuti181, Njuguna200 and biggest of these, the Jonglei Canal on the Upper White Nile, was not Nogrady201; also expeditions, North American and German, from abroad that completed but involved extensive surveys of the Sudd swamp region121a included Cerling40, Hecky114, Kilham142, MacIntyre170, Melack184, 185 and between 1950 and 1980. Vareschi254. These made wide-ranging studies, some of fundamental interest, During 1965 preparations were being made in both Britain and France involving chemical balance87, physical circulation170, primary production184, for intensive team work on two shallow and productive African lakes. That nutrient relationships134 and periodicity of phytoplankton200; also, in Nakuru, on Lake George in Uganda was supported by the Royal Society of London; subsequent trophic transfer to a highly productive food-web255. From Britain, that on Lake Chad by an overseas research organisation (ORSTOM – Green and his associates made contributions for other African water-bodies, Office de la Recherche scientifique et technique outre-mer) of the French including crater lakes in Cameroon94. American and German scientists Government. Work extended over 6–12 years; it was designed to be examined the peculiar vertical stratification in Lake Kivu54, 129, 198, influenced contributory to a global project, the International Biological Programme by heating from below and methane accumulation at depth. (IBP). Organic production was a central theme, with interrelated analyses There was strong interest in the historical reconstruction of lake of planktonic, benthic and fish communities, considered in relation to the environments and biota over very long periods (palaeolimnology). physical and chemical environment. The two lakes were hydrological Evidence of major changes of lake level (some in the rifts exceeding contrasts, with George equable and Chad extremely variable in extent. In 100 m) had come from ancient strand lines199 and mineral deposits81 consequence, there were numerous ecological contrasts. Individual including fossil diatomites. Use was subsequently made of the detailed contributions included work on Lake George by Burgis, Darlington, Dunn, chemical and biological record in layered lake sediments. Pioneer studies Ganf, Greenwood, Gwahaba, McGowan, Moriarty, Tevlin and Viner, and in the 1960s on lakes Victoria, Naivasha and Tanganyika by Livingstone160 on Lake Chad by Bénech, Carmouze, Dejoux, Durand, Duwat, Gac, Gras, and his students Richardson225 and Kendall141 were followed by others in Iltis, Lauzanne, Lemoalle, Lévêque, Loubens, Quensière and Saint-Jean. Ethiopia by Gasse84, 85. In Uganda there was work on Lake George111 and Additional studies were made by visiting scientists. There are extended on swamps by Morrison195 and Hamilton102. Historical reconstruction could summaries of the overall scientific results33, 38, 96 that are major inputs to also be applied to problems of biogeography, as investigated by Dumont tropical freshwater science. and his associates in waters of the Sahel region62, having regard to ‘the Also in the 1960s and 1970s, a number of smaller university-supported Sahara before the desert’ 122 for which French workers had found aquatic projects yielded results of high general interest. That from the University relics – including those of hippopotamus, crocodile and Nile perch – to be of Malawi involved a wide range of studies on Lake Chilwa, another widespread50, 71, 212. hydrologically unstable lake, with events followed over a cycle of drying- Inevitably, many had speculated on the nature of the nutrients most up and re-flooding121, 135. From the University of Khartoum there was an limiting for biological production. Based on chemical surveys, importance exploration of chemical and plankton dynamics, seasonal and inter-annual, was attached to phosphorus, nitrogen246 and sulphur14, 116. More direct in the White Nile and over the river-reservoir cascade system of the Blue evidence was obtained from enrichment bioassays and element uptake; Nile103, 191, 235 where the creation of a new upstream reservoir had examples in East Africa included waters of lakes Victoria69, 72, Naivasha134, repercussions below. From Addis Ababa, Ethiopia, came work on crater 214, Sonachi186 and a fishpond210, in Central Africa various reservoirs226 and lakes222, 265 and rift lakes16, 266; from Elizabethville (later Lumumbashi), Lake Chilwa196, and in West Africa the coastal Ebrié lagoon59. Zaïre, studies of seasonality of reservoir stratification172, river chemistry240 Three projects on major water-bodies were carried out by larger research and phytoplankton production75. A smaller river in Ghana was the base for teams. That on Lake Victoria centred on the understanding and a detailed examination of seasonal progression in benthic invertebrates125, development of fisheries, and applied some novel fishing methods from and one in Gambia for an analysis of solute dynamics153. From Nigeria abroad under the aegis of the FAO127. The new independence of Kenya and there was another seasonal study on a small lake influenced by the annual Tanzania was reflected in the development of two more bases for fisheries harmattan wind regime105, and which extended to day-night vertical and associated research on the lake, at Kisumu203 and Mwanza263. Another migrations of Chaoborus larvae106. In Kenya, further work on time- fisheries-orientated project, on the less known and relatively remote Lake sequences in environments and biota was made on lakes (many of them Turkana in Kenya120, was supported by the Overseas Development Agency soda lakes) of the rift valley – especially Naivasha, Sonachi, Elmenteita of Britain. The third was executed by the equivalent French organisation and Nakuru. This involved university scientists from Nairobi, including ORSTOM and concerned the large, elongate, coastal Ebrié lagoon of the

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Ivory Coast, West Africa. Here there was a complex interplay of inputs nutrient enrichment with enhanced planktonic productivity112, 203 and from sea and freshwater plus pollutants from the adjacent city of Abidjan63. further depletion of oxygen in deep water113. Finally there was, in the These had varied influences on the biota, as shown by studies ranging from 1990s, a troublesome invasion by the water hyacinth, Eichhornia bacteria to fish communities. crassipes. All these events and trends led to further international projects, The period also witnessed a major project of the World Health in which the traditional domination of fisheries and fisheries management Organisation (WHO) in West Africa. In parts of this region a disease was followed by emphasis on ecosystem dynamics and wider socio- (onchocerciasis) was rife due to a parasitic filarial nematode transmitted by economic aspects. There was further attention to water movements as a an insect vector, the blackfly Simulium damnosum complex. The aquatic controlling background factor in lakes, both in Victoria170a and the rift larvae of the blackfly form part of the zoobenthos of rapidly flowing lakes of Malawi209 and Tanganyika45. streams, and were the target of geographically extensive chemical In two countries there was productive cooperation between Swedish and treatment involving the insecticide Abate. Efficacy depended on local scientists supported by the Swedish aid organisation SAREC appropriate ecological knowledge obtained from numerous field studies238. (Department for Research Co-operation, within the Swedish International Further freshwater work was related to the transmission of other parasites Development Agency). For Zimbabwe the focus was the natural economy with a water-connected vector – mosquitoes for malaria91, snails for of Lake Kariba after some 25 years of existence. Local inputs came from schistosomiasis32. the University of Zimbabwe171, 179 and the Kariba Fisheries Institute169. Swedish contributions centred on chemical features158 and aspects of the 1985–2005: further initiatives, large and small planktonic and benthic communities46, 137. With Ethiopia the Swedish contribution was primarily by support to research on rift lakes and crater Special interest continued in the very large, deep and ancient lakes of lakes already under investigation from the University of Addis Ababa139. It Tanganyika and Malawi. Each was the object of a sizeable research group overlapped another overseas cooperation, involving biologists from the over several years; that to Tanganyika was set up from Finland159, that to 188 University of Waterloo in Canada. Lake Awassa was a principal research Malawi from Britain . The results ranged from time-changes of vertical 89, 187 208, 209 1a, 159 164 229 site , but there were also chemical and plankton surveys involving the stratification to food-webs , biodiversity and fish production . comparison of many Ethiopian lakes140, 272 that extended earlier work in the There were, independently, other significant contributions by individual 1960s266. The Austrian Academy of Sciences also gave training in scientists41, 66, many attracted by evolutionary issues raised by the rich 138, 146 45 limnology to numerous African professionals, and supported work on the endemic diversity of cichlid fishes and some invertebrate groups . functional ecology of streams in Kenya180. Also in Kenya there was long- Here Japan and North America are well represented. Further international 164 continued and cooperative study of the rift lake Naivasha, initiated by projects were concerned with biodiversity, cichlid evolution and Harper from Britain. This documented some major changes over time, fisheries. mainly related to water balance and introductions of exotic and in part Lake Victoria, the largest in area of all tropical lakes, was the scene of temperate species107, 108 to the relatively cool lake at moderate altitude. ecological change that had major socio-economic implications. The fishery Wider comparative studies were planned within an international project was greatly altered in the 1980s by the spread of a large predator, the Nile (‘International Decade of the East African Lakes’, IDEAL) that gathered perch (Lates niloticus), after its previous and controversial introduction77, 204 pace in the 1980s and 1990s, and ranged latitudinally from Lake Turkana in the late 1950s. This led to radical change in the food-web and the loss 133 83, 174 to Lake Malawi . These included an emphasis on palaeolimnology, a of previously valued species like Oreochromis esculentus and other 99, 206, 259 263 subject to which others contributed , and which included endemic cichlids . An active project developed on the ecology of consequences of remote and recent climatic change150. haplochromine cichlids, HEST (Haplochromis Ecology Survey Team), In West Africa, as well as in Chad, the French overseas research based on Dutch-Tanzanian cooperation in the southern region of the organisation ORSTOM (later IRD – Institut de Recherche pour le lake263. Its members also contributed more widely on the Victoria fish 264 Développement) continued its work. This included further study of the stocks and fisheries . Although the total fish yield increased after the Ebrié lagoon28, hydrological balance of catchments, lakes and rivers152, 173, spread of Lates, the change disadvantaged most local fishermen and much 35 3 stream faunas and anti-onchocerciasis assessment , colonisation sequence biodiversity was irretrievably lost . Still more radical, and in part related to 98 247 25 of ponds , biomass structure in small water-bodies , and biodiversity of increase in the near-shore human population , was clear evidence of fish communities154, 155.

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Crater lakes continued to be objects of research throughout tropical Africa. From Cameroon there was in August 1986 a most unusual, sudden Acknowledgements and damaging release of carbon dioxide of magmatic origin145; in Ethiopia the long-studied group near Bishoftu (Debre Zeit) showed chemical I am indebted to Geoffrey Fryer, Peter Jackson, Jacques Lemoalle, changes under human influence272; and in western Uganda another group Rosemary Lowe-McConnell and Jean-Jacques Symoens for helpful provided material for the study of within-year changes of stratification and suggestions and criticism. Also to ‘Editions d’IRD’, for permission to use planktonic populations143, 144. Fig. 1 originally published (in ref. 243) by ORSTOM, Paris.

Review and synthesis References Various researchers have chosen to bring together the information available on individual water-bodies. Lakes treated in this way include 1. Abdin, G. (1948). The conditions of growth and periodicity of the Kariba4, 194, Chilwa134, McIlwaine = Chivero248, Chad38, Tanganyika45 and algal flora of the Aswan reservoir (Upper Egypt). Bull. Fac. Sci. Nasser21; rivers include the Nile231 and Niger97. In some cases the material Egypt. Univ. 27, 157–175. surveyed was largely derived from a single extended research project. At 1a. Allison, E.H., Patterson, G., Irvine, K., Thompson, A.B. & Menz, A. the opposite extreme the monograph on the Nile river system, initiated by (1995). The pelagic ecosystem. In: The fishery potential and Rzóska, took in very diverse sources over a large geographical area and productivity of the pelagic zone of Lake Malawi / Niassa (ed. A. historical attribution. Regional surveys included one on West African Menz), pp. 351-367. Natural Resources Institute, Overseas waters132 and another on Ethiopian rift lakes250. African shallow water- Development Administration, UK. bodies were treated in a cooperative international effort with components 2. Balek, J. (1977). Hydrology and water resources in tropical Africa. of directory34, bibliography and synthesis61, 243. Information from some Elsevier, Amsterdam. individual countries, including Sudan60, Ethiopia249, Ghana74, Malawi197 3. Balirwa, J.S., Chapman, C.A., Chapma, L.J., Cowx, I.G., Gebeb, K., and Zimbabwe233, has also been summarised. Kaufman, L., Lowe-McConnell, R., Seehausen, O., Wanink, J.H., Besides the obvious descriptive element, particular water-bodies can Welcomme, R.L. & Witte, F. (2003). Biodiversity and fishery emphasize general topics or themes. For example, lakes Chilwa and Chad sustainability in the Lake Victoria basin: an unexpected marriage? demonstrate time-successions arising from hydrological change, and man- Bioscience 53, 703-715. made lakes like Kariba those from a recent origin. Such general topics 4. Balon, E.K. & Coche, A.G. (1974). Lake Kariba, a man-made have also been treated comparatively from wider experience over tropical tropical ecosystem in Central Africa. Monogr. Biologicae 24. Africa. There are examples for sequences of colonisation183, shallow Junk, The Hague. lakes61, 243, water chemistry246, plankton seasonality109, 114, 242, aquatic 5. Beadle, L.C. (1932a). Scientific results of the Cambridge Expedition vegetation56, floodplain dynamics262, biodiversity of fishes154, 270, fisheries to the East African Lakes, 1930-1. 3. Observations on the research164 and conservation issues25, 165. bionomics of some East African swamps. J. Linn. Soc. Zool. 38, A single book, by Beadle9, has combined (up to 1980) regional 135-155. descriptions and subject reviews over tropical Africa as a whole. Two 6. Beadle, L.C. (1932b). Scientific results of the Cambridge Expedition others211, 245 have taken the still more extended view of general ecological to the East African Lakes, 1930-1. 4. The waters of some East science for tropical inland waters worldwide. Tropical fish communities African Lakes in relation to their fauna and flora. J. Linn. Soc. have been similarly treated163. African experience then provided examples Zool. 38, 157-211. within a generalised scientific framework. This is particularly apt for lake 7. Beadle, L.C. (1957). Respiration in the African swamp worm Alma developments in time, chemical control of aquatic biota, hydrodynamics of emini. J. Exp. Biol. 34, 1-10. large warm lakes, temperature – production relationships, the 8. Beadle, L.C. (1963). Anaerobic life in a tropical crater lake. Nature, determination of seasonality at low latitudes, and the localisation or Lond. 200, 1223-1224. adaptive radiation of genetic novelty. 9. Beadle, L.C. (1981). The inland waters of tropical Africa. An introduction to tropical limnology. Second edition. Longman, London.

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